Editing Santalales

{{Short description|Order of flowering plants}}
{{Automatic taxobox
|taxon = Santalales
|authority = R.Br. ex Bercht. & J.Presl<ref name=WFO/>
|image = Starr_041006-0222_Santalum_haleakalae.jpg
|image_caption = ''[[Santalum haleakalae]]''
|subdivision_ranks = Families
|subdivision = See text
|synonyms = Balanophorales
}}

The '''Santalales''' are an [[order (biology)|order]] of [[flowering plant]]s in the [[dicotyledon]]s. Well-known members of the Santalales include [[Santalum|sandalwoods]] and the many species of [[mistletoe]]s. The order has a [[cosmopolitan distribution]], but is heavily concentrated in tropical and subtropical regions. It derives its name from its [[type genus]], ''[[Santalum]]'' (sandalwood).

== Overview ==
Many of the members of the order are [[parasitic plant]]s, mostly hemiparasites, able to produce sugars through [[photosynthesis]], but tapping the stems or roots of other plants to obtain water and minerals; some (e.g. ''[[Arceuthobium]]'') are obligate [[parasite]]s, have low concentrations of [[chlorophyll]] within their shoots (1/5 to 1/10 of that found in their host's foliage), and derive the majority of their sustenance from their hosts' vascular tissues (water, [[micronutrient|micro-]] and [[macronutrient]]s, and sucrose).

Most have [[seed]]s without [[testa (botany)|testa]]e (seed coats), which is unusual for flowering plants.

== Classification ==
The [[APG IV system]] of 2016 includes seven families.<ref name=APGIV/> As in the earlier [[APG III system]], it was accepted that Olacaceae ''sensu lato'' was [[Paraphyly|paraphyletic]] but new family limits were not proposed as relationships were considered uncertain.<ref name=APGIII/><ref name=APGIV/> {{As of|2021|July}}, this seven-family division of the Santalales was explicitly accepted by the [[World Flora Online]],<ref name=WFO/> and implicitly by [[Plants of the World Online]], in that it accepted none of the extra families recognized by other sources. The seven families are:
* [[Balanophoraceae]]
* [[Loranthaceae]]
* [[Misodendraceae]]
* [[Olacaceae]]
* [[Opiliaceae]]
* [[Santalaceae]]
* [[Schoepfiaceae]]

When only these families are recognized, one possible phylogenetic relationship among them is shown below. Support for some of the nodes is weak,<ref name=APweb/> and at least two families, Olacaceae ''[[Sensu|s.l.]]'' and Balanophoraceae ''s.l.'', are not monophyletic:<ref name=APweb/><ref name=Nick20/> 
{{clade
|label1=Santalales
|1={{clade
   |1=[[Olacaceae]] ''s.l.''|state1=double
   |2={{clade
      |1=[[Balanophoraceae]] ''s.l.''|state1=double
      |2={{clade
         |1={{clade
            |1=[[Loranthaceae]]
            |2={{clade
               |1=[[Misodendraceae]]
               |2=[[Schoepfiaceae]]
               }}
            }}
         |2={{clade
            |1=[[Opiliaceae]]
            |2=[[Santalaceae]] ''s.l.''
            }}
         }}
      }}
   }}
}}

A summary of the circumscription and phylogeny of the Santalales published in 2020 used 20 rather than seven families. Olacaceae ''s.l.'' was divided into seven families, Balanophoraceae ''s.l.'' was divided into two, and Santalaceae ''s.l.'' into seven.<ref name=Nick20/> {{As of|2021|July}}, the [[Angiosperm Phylogeny Website]] accepted the families resulting from the division of Olacaceae ''s.l.'' and Balanophoraceae ''s.l.'' but not those from the division of Santalaceae ''s.l.''<ref name=APweb/>
{| class="wikitable"
|+ Division of the ''sensu lato'' Santalales families according to Nickrent (2020)<ref name=Nick20/>
|-style="vertical-align: top;"
|
*Olacaceae ''s.l.''
**Aptandraceae
**Coulaceae
**Erythropalaceae
**Octoknemaceae
**Olacaceae ''s.s.''
**Strombosiaceae
**Ximeniaceae 
|
*Balanophoraceae ''s.l.''
**Balanophoraceae ''s.s.''
**Mystropetalaceae
|
*Santalaceae ''s.l.''
**Amphorogynaceae
**Cervantesiaceae
**Comandraceae
**Nanodeaceae
**Santalaceae ''s.s.''
**Thesiaceae
**Viscaceae
|}

===Earlier systems===
In the [[Dahlgren system|classification system]] of [[Rolf Dahlgren|Dahlgren]], the Santalales were in the [[superorder]] Santaliflorae (also called Santalanae). The [[Cronquist system]] (1981) used this circumscription:<ref name=Cron81/>
* order Santalales
:* family Medusandraceae – sole genus ''[[Medusandra]]'' now in family [[Peridiscaceae]],<ref name="POWO_26132-1">{{cite web |title=''Medusandra'' Brenan |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:26132-1 |access-date=2021-07-20 }}</ref> order [[Saxifragales]]<ref name=APGIV/>
:* family [[Dipentodontaceae]] – now in order [[Huerteales]] in APG IV<ref name=APGIV/>
:* family [[Olacaceae]]
:* family [[Opiliaceae]]
:* family [[Santalaceae]]
:* family [[Misodendraceae]]
:* family [[Loranthaceae]]
:* family [[Viscaceae]] - sunk into Santalaceae ''s.l.'' in the seven-family system
:* family [[Eremolepidaceae]] - sunk into Santalaceae ''s.l.'' in the seven-family system
:* family [[Balanophoraceae]]

==Gallery of type genera==
<gallery mode=packed>
Balanophora indica2.jpg |alt=flowers |''[[Balanophora fungosa]]'' 
Loranthus europaeus (Riemenmistel) IMG 9783.jpg| alt=flowers and foliage |''[[Loranthus europaeus]]'' 
Misodendrum punctulatum.jpg |alt=flowers and foliage |''[[Misodendrum punctulatum]]''
Olax imbricata Roxb. (8503288374).jpg |alt=flowers and foliage |''[[Olax imbricata]]''
Opilia amentacea.jpg |alt=botanical illustration |''[[Opilia amentacea]]''
Santalum album - Indian Botanic Garden - Howrah 2013-03-31 5743.JPG |alt=foliage |''[[Santalum album]]''
Schoepfia brasiliensis A.DC. - Flickr - Alex Popovkin, Bahia, Brazil (11).jpg |alt=foliage |''[[Schoepfia brasiliensis]]''
</gallery>

==References==
{{Commons category}}
{{Wikispecies|santalales}}
{{Reflist|refs=
<ref name=APGIII>{{Cite journal |last=Angiosperm Phylogeny Group |year = 2009 |title = An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal = Botanical Journal of the Linnean Society |volume = 161 |issue = 2 |pages = 105–121 |doi = 10.1111/j.1095-8339.2009.00996.x |doi-access = free |hdl = 10654/18083 |hdl-access = free }}</ref>

<ref name=APGIV>{{Citation |mode=cs1 |last=Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |doi-access=free }}</ref>

<ref name=APweb>{{Cite web |last=Stevens |first=P.F. |title=Santalales |work=Angiosperm Phylogeny Website |url=http://www.mobot.org/MOBOT/research/APweb/orders/santalalesweb2.htm#Santalales |access-date=2021-07-20 }}</ref>

<ref name=Cron81>{{cite book |last=Cronquist |first=A. |year=1981 |title=An integrated system of classification of flowering plants |publisher=[[Columbia University Press]] |location=New York |url=https://archive.org/details/integratedsystem0000cron |url-access=registration |isbn=978-0-231-03880-5}}</ref>

<ref name=Nick20>{{Citation |mode=cs1 |last1=Nickrent |first1=Daniel L. |date=2020 |title=Parasitic angiosperms: How often and how many? |journal=Taxon |volume=69 |issue=1 |pages=5–27 |doi=10.1002/tax.12195 }}</ref>

<ref name=WFO>{{Citation |mode=cs1 |title=Santalales R.Br. ex Bercht. & J.Presl |website=World Flora Online |url=http://www.worldfloraonline.org/taxon/wfo-9000000477 |access-date=2021-07-20 }}</ref>
}}
* {{cite book|last = Hawksworth|first = FG|title = Dwarf mistletoes : biology, pathology, and systematics |publisher = USDA For. Serv. Agric. Handb|year = 1996|pages = 409}}
* {{cite book|last1 = Soltis|first1 = Douglas E. |author-link1=Douglas E. Soltis |last2=Soltis |first2=Pamela S. |author-link2 = Pamela S. Soltis |first3=Peter K. |last3=Endress |first4=Mark W. |last4=Chase |author-link4=Mark Wayne Chase |title = Phylogeny & Evolution of Angiosperms |publisher = Sinauer Associates |date = 2005-06-15|pages = 370|isbn = 978-0-87893-817-9}}
* [https://parasiticplants.siu.edu/ListParasites.html#anchor530678 Santalales on the Parasitic Plant Connection web page]
* [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Undef&id=41947&lvl=3&p=mapview&p=has_linkout&p=blast_url&p=genome_blast&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]

{{Angiosperm orders}}
{{Taxonbar|from=Q21851}}
{{Authority control}}

[[Category:Santalales| ]]
[[Category:Superasterids]]
[[Category:Angiosperm orders]]